Christopher A. Pearl

Fire and amphibians in North America

Information on amphibian responses to fire and fuel reduction practices is critically needed due to potential declines of species and the prevalence of new, more intensive fire management practices in North American forests. The goals of this review are to summarize the known and potential effects of fire and fuels management on amphibians and their aquatic habitats, and to identify information gaps to help direct future scientific research. Amphibians as a group are taxonomically and ecologically diverse; in turn, responses to fire and associated habitat alteration are expected to vary widely among species and among geographic regions. Available data suggest that amphibian responses to fire are spatially and temporally variable and incompletely understood. Much of the limited research has addressed short-term (1‐3 years) effects of prescribed fire on terrestrial life stages of amphibians in the southeastern United States. Information on the long-term negative effects of fire on amphibians and the importance of fire for maintaining amphibian communities is sparse for the majority of taxa in North America. Given the size and severity of recent wildland fires and the national effort to reduce fuels on federal lands, future studies are needed to examine the effects of these landscape disturbances on amphibians. We encourage studies to address population-level responses of amphibians to fire by examining how different life stages are affected by changes in aquatic, riparian, and upland habitats. Research designs need to be credible and provide information that is relevant for fire managers and those responsible for assessing the potential effects of various fuel reduction alternatives on rare, sensitive, and endangered amphibian species. # 2003 Elsevier Science B.V. All rights reserved.

Asymmetrical effects of introduced Bullfrogs (Rana catesbeiana) on native ranid frogs in Oregon

Abstract Introduced American Bullfrogs (Rana catesbeiana) have become widely established in the Pacific Northwest over the last century and are thought to be an important predator of native amphibians throughout the western United States. The Northern Red-Legged Frog (Rana aurora aurora) and Oregon Spotted Frog (Rana pretiosa) historically coexisted in portions of the Pacific Northwest now invaded by R. catesbeiana, but R. pretiosa has declined more severely than R. a. aurora. We investigated whether microhabitat and behavioral differences that facilitate sympatric coexistence of the natives predict which species is more susceptible to predation by introduced R. catesbeiana. Our laboratory experiments demonstrate that R. catesbeiana adults prefer aquatic microhabitats, that R. pretiosa juveniles are more aquatic than R. a. aurora, and that adult R. catesbeiana consume more R. pretiosa than R. a. aurora juveniles. Mean and maximum jump distances of R. pretiosa were shorter than equally sized R. a. aurora, and the difference between these two species increased with larger frog sizes. Our examination of field survey data indicates that R. pretiosa coexist with R. catesbeiana less frequently than R. a. aurora. We conclude that R. catesbeiana is a greater threat to survival of R. pretiosa than to R. a. aurora and suggest that microhabitat use and escape abilities of native ranid frogs may be linked to this asymmetrical effect. Analysis of behavioral and microhabitat differences among related native species may be a useful tool in predicting the effects of introduced predators on amphibians and can assist in developing conservation priorities for these species.

Behavioral responses of anuran larvae to chemical cues of native and introduced predators in the Pacific Northwestern United States

Abstract We compared behavioral responses of larvae of three Pacific Northwest anurans from different hydroperiods to water borne cues of native and introduced predators. Two native anurans (Pacific Treefrog, Pseudacris regilla, and Northern Red-Legged Frog, Rana aurora aurora) and introduced Bullfrogs (Rana catesbeiana) responded to water conditioned by native Redside Shiners (Richardsonius balteatus) by increasing refuge use. The larvae of the two native anurans differed in their response to introduced predator cues. Rana aurora aurora, which occur in temporary and permanent waters, responded to both introduced Bluegill Sunfish (Lepomis macrochirus) and introduced Crayfish (Procambarus clarkii). Pseudacris regilla, which occur primarily in temporary ponds, did not respond to water borne cues from either introduced predator. The broader responses of R. a. aurora may indicate greater behavioral plasticity or more exposure to novel predators than experienced by P. regilla. Larvae of introduced R. catesbeiana responded strongly to cues from two fish native to the Pacific northwest but did not alter behavior in response to any of five potential predators with which they coexist in their native range. Fish that occur with R. catesbeiana in their native range generally find Bullfrog larvae unpalatable. This pattern suggests that Bullfrog larvae can recognize cues of novel predators that may find them palatable, which could contribute to their success as an invasive species in the region.

AMPHIBIAN OCCURRENCE AND AQUATIC INVADERS IN A CHANGING LANDSCAPE: IMPLICATIONS FOR WETLAND MITIGATION IN THE WILLAMETTE VALLEY, OREGON, USA

Despite concern about the conservation status of amphibians in western North America, few field studies have documented occurrence patterns of amphibians relative to potential stressors. We surveyed wetland fauna in Oregon’s Willamette Valley and used an information theoretic approach (AIC) to rank the associations between native amphibian breeding occurrence and wetland characteristics, non-native aquatic predators, and landscape characteristics in a mixed urban-agricultural landscape. Best predictors varied among the five native amphibians and were generally consistent with life history differences. Pacific tree frog (Pseudacris regilla) and long-toed salamander (Ambystoma macrodactylum) occurrence was best predicted by the absence of non-native fish. Northern red-legged frog (Rana a. aurora) and northwestern salamander (Ambystoma gracile) were most strongly related to wetland vegetative characteristics. The occurrence habitats, was best predicted by greater forest cover within 1 km. The absence of non-native fish was a strong predictor of occurrence for four of the five native species. In contrast, amphibians were not strongly related to native fish presence. We found little evidence supporting negative effects of the presence of breeding populations of bullfrog (Rana catesbeiana) on any native species. Only the two Ambystoma salamanders were associated with wetland permanence. Northwestern salamanders (which usually have a multi-year larval stage) were associated with permanent waters, while long-toed salamanders were associated with temporary wetlands. Although all the species make some use of upland habitats, only one (rough-skinned newt) was strongly associated with surrounding landscape conditions. Instead, our analysis suggests that within-wetland characteristics best predict amphibian occurrence in this region. We recommend that wetland preservation and mitigation efforts concentrate on sites lacking non-native fish for the conservation of native amphibians in the Willamette Valley and other western lowlands.

OPTICAL CHARACTERISTICS OF NATURAL WATERS PROTECT AMPHIBIANS FROM UV-B IN THE U.S. PACIFIC NORTHWEST

Increased exposure to ultraviolet-B (UV-B) radiation has been proposed as a major environmental stressor leading to global amphibian declines. Prior experimental evidence from the U.S. Pacific Northwest (PNW) indicating the acute embryonic sensitivity of at least four amphibian species to UV-B has been central to the literature about amphibian decline. However, these results have not been expanded to address population-scale effects and natural landscape variation in UV-B transparency of water at amphibian breeding sites: both necessary links to assess the importance of UV-B for amphibian declines. We quantified the UV-B transparency of 136 potential amphibian breeding sites to establish the pattern of UV-B exposure across two montane regions in the PNW. Our data suggest that 85% of sites are naturally protected by dissolved organic matter in pond water, and that only a fraction of breeding sites are expected to experience UV-B intensities exceeding levels associated with elevated egg mortality. Thus, the spectral characteristics of natural waters likely mediate the physiological effects of UV-B on amphibian eggs in all but the clearest waters. These data imply that UV-B is unlikely to cause broad amphibian declines across the landscape of the American Northwest.

Occurrence of the Amphibian Pathogen Batrachochytrium Dendrobatidis in the Pacific Northwest

Abstract Chytridiomycosis (infection by the fungus Batrachochytrium dendrobatidis) has been associated with amphibian declines in at least four continents. We report results of disease screens from 210 pond-breeding amphibians from 37 field sites in Oregon and Washington. We detected B. dendrobatidis on 28% of sampled amphibians, and we found ≥ 1 detection of B. dendrobatidis from 43% of sites. Four of seven species tested positive for B. dendrobatidis, including the Northern Red-Legged Frog (Rana aurora), Columbia Spotted Frog (Rana luteiventris), and Oregon Spotted Frog (Rana pretiosa). We also detected B. dendrobatidis in nonnative American Bullfrogs (Rana catesbeiana) from six sites in western and central Oregon. Our study and other recently published findings suggest that B. dendrobatidis has few geographic and host taxa limitations among North American anurans. Further research on virulence, transmissibility, persistence, and interactions with other stressors is needed to assess the potential impact of B. dendrobatidis on Pacific Northwestern amphibians.

Using occupancy models to understand the distribution of an amphibian pathogen, Batrachochytrium dendrobatidis

Batrachochytrium dendrobatidis is a fungal pathogen that is receiving attention around the world for its role in amphibian declines. Study of its occurrence patterns is hampered by false negatives: the failure to detect the pathogen when it is present. Occupancy models are a useful but currently underutilized tool for analyzing detection data when the probability of detecting a species is <1. We use occupancy models to evaluate hypotheses concerning the occurrence and prevalence of B. dendrobatidis and discuss how this application differs from a conventional occupancy approach. We found that the probability of detecting the pathogen, conditional on presence of the pathogen in the anuran population, was related to amphibian development stage, day of the year, elevation, and human activities. Batrachochytrium dendrobatidis was found throughout our study area but was only estimated to occur in 53.4% of 78 populations of native amphibians and 66.4% of 40 populations of nonnative Rana catesbeiana tested. We found little evidence to support any spatial hypotheses concerning the probability that the pathogen occurs in a population, but did find evidence of some taxonomic variation. We discuss the interpretation of occupancy model parameters, when, unlike a conventional occupancy application, the number of potential samples or observations is finite.

Problems and opportunities managing invasive bullfrogs: is there any hope?

INTRODUCTIONThe American Bullfrog (Rana catesbeiana Shaw) is a widely introduced andinvasive anuran that is frequently blamed for population declines of indigenousspecies (Bury and Whelan 1984). Once established, Bullfrog populationsare often either difficult or impossible to eradicate depending on habitat andlandscape features (Schwalbe and Rosen 1988, Doubledee et al. 2003,Govindarajulu et al. 2005). Bullfrogs are representative of a large but neglectedsuite of non-indigenous species (NIS) that are characterized by: (1) a broadinvasion that is well established in some areas; (2) a lack of obvious economicimpacts compared to some other invasive species; and (3) a lack of reasonablyfeasible control methods. Despite demonstrated conservation concerns, invasivespeciesliketheBullfrogdonottendtoattracttheresourcesnecessarytoattemptlarge scale management because of their lack of economic impact and thedifficulty of control methods. This leaves biologists responsible for managinghabitats invaded by such species with little hope and few options for promotingthe persistence of sensitive indigenous species. With these issues in mind, weconsider the case of the Bullfrog, review management options, and suggestdirections for future research with this and similar species.Bullfrogs are among the most successful vertebrate invaders and are con-sidered by the IUCN Invasive Species Specialist Group to be among the 100worst invaders in the world (http://www.issg.org/database/welcome). The679

Observations of Interspecific Amplexus Between Western North American Ranid Frogs and the Introduced American Bullfrog (Rana catesbeiana) and an Hypothesis Concerning Breeding Interference

Abstract Introduced American bullfrogs (Rana catesbeiana) come in contact with native amphibians on four continents and are well established in lowlands of western North America. To date, research on the effects of introduced bullfrogs on native frogs has focused on competition and predation, and is based largely on larval interactions. We present observations of interspecific amplexus between bullfrogs and two native ranid frogs (R. aurora and R. pretiosa) from six sites across the Pacific Northwest that imply that this interaction is more widespread than currently recognized. Our observations indicate that R. catesbeiana juveniles and subadults in this region are of appropriate size to elicit marked amplectic responses from males of both native species. Our literature review suggests that greater opportunity may exist for pairings between R. catesbeiana and native R. aurora or R. pretiosa than among syntopic native ranids in western North America. We hypothesize that interspecific amplexus with introduced R. catesbeiana could result in reproductive interference with negative demographic consequences in native ranid populations that have been reduced or altered by other stressors.

Low Prevalence of Chytrid Fungus (Batrachochytrium dendrobatidis) in Amphibians of U.S. Headwater Streams

Abstract Many declines of amphibian populations have been associated with chytridiomycosis, a disease caused by the aquatic fungus Batrachochytrium dendrobatidis (Bd). Despite the relatively high prevalence of chytridiomycosis in stream amphibians globally, most surveys in North America have focused primarily on wetland-associated species, which are frequently infected. To better understand the distribution and prevalence of Bd in headwater amphibian communities, we sampled 452 tailed frogs (Ascaphus truei and Ascaphus montanus) and 304 stream salamanders (seven species in the Dicamptodontidae and Plethodontidae) for Bd in 38, first- to third-order streams in five montane areas across the United States. We tested for presence of Bd by using PCR on skin swabs from salamanders and metamorphosed tailed frogs or the oral disc of frog larvae. We detected Bd on only seven individuals (0.93%) in four streams. Based on our study and results from five other studies that have sampled headwater- or seep-associated amphibians in the United States, Bd has been detected on only 3% of 1,322 individuals from 21 species. These results differ strongly from surveys in Central America and Australia, where Bd is more prevalent on stream-breeding species, as well as results from wetland-associated anurans in the same regions of the United States that we sampled. Differences in the prevalence of Bd between stream- and wetland-associated amphibians in the United States may be related to species-specific variation in susceptibility to chytridiomycosis or habitat differences.